Determinants of fine-scale plant diversity in dry calcareous grasslands within the Baltic Sea region

We used an extensive dataset (1220 vegetation plots of 1 m(2)) to study vegetation gradients and fine-scale plant diversity in dry calcareous grasslands (including alvar grasslands) in the Baltic Sea region. The study area covers the entire European distributional range of alvar habitats: Sweden (Oland, Gotland, Gotaland), Estonia (Saaremaa, Hiiumaa, north Estonia, west Estonia), and western Russia (Izhora, lzborsk). Fine-scale plant diversity was characterized by species richness and standardized phylogenetic diversity (comparing the observed mean pairwise phylogenetic distance (MPD) with MPD values from random communities). Ordination techniques (DCA) were used to characterize the main vegetation gradient. Variables describing local environment, climate, the biogeographic composition of the plant communities, and geographic location were related to fine-scale species richness and phylogenetic diversity using variation partitioning techniques and linear mixed models. The main vegetation gradient in the dry calcareous grasslands in the Baltic Sea region had a strong geographic component, was associated with soil depth, species' stress- and disturbance-tolerance and the age of the grassland habitat. Fine-scale phylogenetic diversity and species richness were negatively associated suggesting that these two diversity components are influenced by different sets of environmental and historical parameters. Fine-scale species richness was unimodally associated with the main vegetation gradient, and the highest levels of species richness were found under intermediate environmental (disturbance, light conditions and temperature) conditions where there was a mixture of species from different biogeographic regions. In contrast to species richness, fine-scale phylogenetic diversity was negatively associated with the main vegetation gradient. The highest phylogenetic diversity was found in the extremely thin-soiled alvar grasslands in Gotaland and on the Baltic islands (especially on Oland) where the high phylogenetic diversity is likely to be a reflection of a long history of continuous openness that has allowed time for the "collection" of phylogenetically different species within these unique habitats. (C) 2012 Elsevier B.V. All rights reserved